{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T13:37:54Z","timestamp":1791466674813,"version":"4.3.4"},"reference-count":70,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2025,12,11]],"date-time":"2025-12-11T00:00:00Z","timestamp":1765411200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Vet. Sci."],"abstract":"<jats:p>\n                    Paratuberculosis (PTB) is a chronic enteritis caused by\n                    <jats:italic>Mycobacterium avium<\/jats:italic>\n                    subsp.\n                    <jats:italic>paratuberculosis<\/jats:italic>\n                    (Map). Genome-wide association studies revealed a significant enrichment of the keratinization pathway in cattle with multifocal lesions, suggesting a potential role of cytokeratins (CKs) in PTB resilience mechanisms. To confirm this, the amount of CK-expressing cells was analyzed in granulomas from the distal jejunum (DJE) and jejunal lymph nodes (DJELN) of animals with focal, multifocal, and diffuse lesions, and in control animals without lesions. Quantitative double-immunohistochemical (D-IHC) analysis [using Iba1 (ionized calcium-binding adapter molecule-1) and CK, as specific markers of macrophages and epithelial cells, respectively] showed that animals with multifocal lesions had the highest numbers of double-Iba1\/CK positive cells [epithelioid macrophages (EMs)]. Significant differences were observed with the focal (\n                    <jats:italic>p<\/jats:italic>\n                    \u202f&amp;lt;\u202f0.001), diffuse (\n                    <jats:italic>p<\/jats:italic>\n                    \u202f=\u202f0.038), and control groups (\n                    <jats:italic>p<\/jats:italic>\n                    \u202f&amp;lt;\u202f0.001) in JELN. Similarly, these animals showed higher numbers of single-CK expressing cells in JELN and DJE. Two EM distribution patterns were observed. In Pattern 1, mostly observed in animals with multifocal lesions (low Map load, no clinical signs), EMs form a barrier-like arrangement around the granuloma, while in Pattern 2, mainly found in animals with diffuse lesions (high Map load, clinical signs), EMs have a diffuse arrangement throughout the granuloma. These findings suggest that animals with multifocal lesions might represent a resilient phenotype that controls Map infection and disease progression. This is achieved through the formation of ordered granulomas that prevent Map dissemination and maintain tissue integrity, which are characteristic features of resilient animals. CK could be considered a potential biomarker of PTB resilience.\n                  <\/jats:p>","DOI":"10.3389\/fvets.2025.1690841","type":"journal-article","created":{"date-parts":[[2025,12,11]],"date-time":"2025-12-11T15:29:37Z","timestamp":1765466977000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":1,"title":["Cytokeratin expression and distribution pattern of epithelioid macrophages in animals with different pathological forms of bovine paratuberculosis: potential role in resilience to PTB"],"prefix":"10.3389","volume":"12","author":[{"given":"Alejandra Isabel","family":"Navarro Le\u00f3n","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marta","family":"Mu\u00f1oz","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cristina","family":"Blanco-V\u00e1zquez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Natalia","family":"Iglesias","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tania","family":"Iglesias","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mar\u00eda","family":"Canive","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gerard","family":"Badia-Bringu\u00e9","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marta","family":"Alonso-Hearn","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ana","family":"Balseiro","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rosa","family":"Casais","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2025,12,11]]},"reference":[{"key":"ref1","doi-asserted-by":"publisher","first-page":"607","DOI":"10.1016\/S1473-3099(07)70211-6","article-title":"Mycobacterium 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culture for purpose-specific testing for paratuberculosis","volume":"89","author":"Nielsen","year":"2006","journal-title":"J Dairy Sci"},{"key":"ref8","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1016\/j.jcpa.2004.04.004","article-title":"Characterization of macrophages and occurrence of T cells in intestinal lesions of subclinical paratuberculosis in goats","volume":"131","author":"Valheim","year":"2004","journal-title":"J Comp Pathol"},{"key":"ref9","doi-asserted-by":"publisher","first-page":"723","DOI":"10.1136\/jcp.36.7.723","article-title":"Granulomatous inflammation \u2013 a review","volume":"36","author":"Williams","year":"1983","journal-title":"J Clin Pathol"},{"key":"ref10","article-title":"Granuloma","volume-title":"StatPearls (internet)","author":"Williams","year":"2022"},{"key":"ref11","doi-asserted-by":"publisher","first-page":"184","DOI":"10.1016\/j.jcpa.2005.04.007","article-title":"Histopathological classification of lesions associated with natural 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granulomas","volume":"36","author":"Wanat","year":"2014","journal-title":"Am J Dermatopathol"},{"key":"ref23","doi-asserted-by":"publisher","first-page":"361","DOI":"10.1016\/0092-8674(82)90234-3","article-title":"Correlation of specific keratins with different types of epithelial differentiation, monoclonal antibody studies","volume":"30","author":"Tseng","year":"1982","journal-title":"Cell"},{"key":"ref24","doi-asserted-by":"publisher","first-page":"e0121625","DOI":"10.1371\/journal.pone.0121625","article-title":"Expression of cellular components in granulomatous inflammatory response in Piaractus mesopotamicus model","volume":"10","author":"Manrique","year":"2015","journal-title":"PLoS One"},{"key":"ref25","doi-asserted-by":"publisher","first-page":"429","DOI":"10.1177\/030098588902600508","article-title":"Chronic inflammatory cells with epithelial cell characteristics in teleost fishes","volume":"26","author":"Noga","year":"1989","journal-title":"Vet 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Ther"},{"key":"ref29","doi-asserted-by":"publisher","first-page":"20177","DOI":"10.1038\/s41598-021-99672-4","article-title":"Identification of loci associated with pathological outcomes in Holstein cattle infected with Mycobacterium avium subsp. paratuberculosis using whole-genome sequence data","volume":"11","author":"Canive","year":"2021","journal-title":"Sci Rep"},{"key":"ref30","doi-asserted-by":"publisher","first-page":"600","DOI":"10.1016\/j.ceb.2013.06.008","article-title":"Networking galore, intermediate filaments and cell migration","volume":"25","author":"Chung","year":"2013","journal-title":"Curr Opin Cell Biol"},{"key":"ref31","doi-asserted-by":"publisher","first-page":"616","DOI":"10.1038\/nature13903","article-title":"P63(+) Krt5(+) distal airway stem cells are essential for lung 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Immunol"},{"key":"ref35","doi-asserted-by":"publisher","first-page":"936","DOI":"10.1126\/science.1214935","article-title":"Disease tolerance as a defense strategy","volume":"335","author":"Medzhitov","year":"2012","journal-title":"Science"},{"key":"ref36","doi-asserted-by":"publisher","first-page":"3346","DOI":"10.2527\/jas.2017.1479","article-title":"Selection on resilience improves disease resistance and tolerance to infections","volume":"95","author":"Mulder","year":"2017","journal-title":"J Anim Sci"},{"key":"ref37","doi-asserted-by":"publisher","first-page":"483","DOI":"10.1016\/j.it.2014.08.001","article-title":"Tissue damage control in disease tolerance","volume":"35","author":"Soares","year":"2014","journal-title":"Trends Immunol"},{"key":"ref38","doi-asserted-by":"publisher","first-page":"328","DOI":"10.1186\/s12917-017-1254-6","article-title":"Case definition terminology for paratuberculosis (Johne's disease)","volume":"13","author":"Whittington","year":"2017","journal-title":"BMC Vet Res"},{"key":"ref39","doi-asserted-by":"publisher","first-page":"e0236336","DOI":"10.1371\/journal.pone.0236336","article-title":"Detection of latent forms of Mycobacterium avium subsp. paratuberculosis infection using host biomarker-based ELISAs greatly improves paratuberculosis diagnostic sensitivity","volume":"15","author":"Blanco-V\u00e1zquez","year":"2020","journal-title":"PLoS One"},{"key":"ref40","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1053\/jcpa.2000.0430","article-title":"Expression of cytokeratins and vimentin in normal and neoplastic tissue from the bovine female reproductive tract","volume":"124","author":"P\u00e9rez-Mart\u00ednez","year":"2001","journal-title":"J Comp Pathol"},{"key":"ref41","doi-asserted-by":"publisher","first-page":"459","DOI":"10.1002\/mrd.1080410408","article-title":"Distribution of the intermediate filament proteins vimentin, keratin, and desmin in the bovine ovary","volume":"41","author":"Van den Hurk","year":"1995","journal-title":"Mol Reprod Dev"},{"key":"ref42","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1016\/j.jcpa.2014.07.003","article-title":"Immunohistochemical expression of ionized calcium binding adapter molecule 1 in cutaneous histiocytic proliferative, neoplastic and inflammatory disorders of dogs and cats","volume":"151","author":"Pierezan","year":"2014","journal-title":"J Comp Pathol"},{"key":"ref43","doi-asserted-by":"publisher","first-page":"1424","DOI":"10.1177\/27.11.92498","article-title":"The unlabeled antibody method. Contrasting color staining of paired pituitary hormones without antibody removal","volume":"27","author":"Sternberger","year":"1979","journal-title":"J Histochem Cytochem"},{"key":"ref44","doi-asserted-by":"publisher","first-page":"981","DOI":"10.1177\/29.8.7024402","article-title":"Inhibition of endogenous tissue alkaline phosphatase with the use of alkaline phosphatase conjugates in immunohistochemistry","volume":"29","author":"Ponder","year":"1981","journal-title":"J Histochem Cytochem"},{"key":"ref45","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1038\/mi.2010.66","article-title":"Intestinal macrophages and response to microbial encroachment","volume":"4","author":"Smith","year":"2011","journal-title":"Mucosal Immunol"},{"key":"ref46","first-page":"153","article-title":"Experimental pine pollen granulomatous pneumonia in the rat","volume":"49","author":"Adams","year":"1966","journal-title":"Am J Pathol"},{"key":"ref47","doi-asserted-by":"publisher","first-page":"154","DOI":"10.1016\/j.vetmic.2008.02.017","article-title":"Association of severity of enteric granulomatous inflammation with disseminated Mycobacterium avium subspecies paratuberculosis infection and antemortem test results for paratuberculosis in dairy cows","volume":"131","author":"Dennis","year":"2008","journal-title":"Vet Microbiol"},{"key":"ref48","doi-asserted-by":"publisher","first-page":"378","DOI":"10.1354\/vp.41-4-378","article-title":"Sequential development of histologic lesions and their relationship with bacterial isolation, fecal shedding, and immune responses during progressive stages of experimental infection of lambs with Mycobacterium avium subsp. paratuberculosis","volume":"41","author":"Kurade","year":"2004","journal-title":"Vet Pathol"},{"key":"ref49","doi-asserted-by":"publisher","first-page":"163","DOI":"10.1016\/s0065-2776(08)60443-5","article-title":"The structure and function of monocytes and macrophages","volume":"9","author":"Cohn","year":"1968","journal-title":"Adv Immunol"},{"key":"ref50","doi-asserted-by":"publisher","first-page":"187","DOI":"10.1002\/path.1711050305","article-title":"The origin, properties and fate of epithelioid cells","volume":"105","author":"Papadimitriou","year":"1971","journal-title":"J Pathol"},{"key":"ref51","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1002\/path.1711060104","article-title":"The ultrastructure of high- and low-turnover inflammatory granulomata","volume":"106","author":"Papadimitriou","year":"1972","journal-title":"J Pathol"},{"key":"ref52","doi-asserted-by":"publisher","first-page":"7820","DOI":"10.1128\/IAI.69.12.7820-7831.2001","article-title":"Dynamic nature of host-pathogen interactions in Mycobacterium marinum granulomas","volume":"69","author":"Bouley","year":"2001","journal-title":"Infect Immun"},{"key":"ref53","doi-asserted-by":"publisher","first-page":"986","DOI":"10.1038\/ni.2705","article-title":"Tissue-resident macrophages","volume":"14","author":"Davies","year":"2013","journal-title":"Nat Immunol"},{"key":"ref54","doi-asserted-by":"publisher","first-page":"502","DOI":"10.1016\/j.immuni.2009.06.025","article-title":"Intestinal lamina propria dendritic cell subsets have different origins and functions","volume":"31","author":"Varol","year":"2009","journal-title":"Immunity"},{"key":"ref55","doi-asserted-by":"publisher","first-page":"352","DOI":"10.1038\/nri3211","article-title":"Revisiting the role of the granuloma in tuberculosis","volume":"12","author":"Ramakrishnan","year":"2012","journal-title":"Nat Rev Immunol"},{"key":"ref56","doi-asserted-by":"publisher","first-page":"693","DOI":"10.1016\/s1074-7613(02)00475-2","article-title":"Real-time visualization of mycobacterium-macrophage interactions leading to initiation of granuloma formation in zebrafish 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